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Ecology Chemical Ecology
by
André Kessler

Introduction

“Ours is a world of sights and sounds. We live by our eyes and ears and tend generally to be oblivious to the chemical happenings in our surrounds. Such happenings are ubiquitous. All organisms engender chemical signals, and all, in their respective ways, respond to the chemical emissions of others. The result is a vast communicative interplay, fundamental to the fabric of life” (Eisner and Meinwald 1995, p. v), cited under General Overviews). Chemical ecology is the study of ecological interactions between organisms mediated by chemicals produced by those organisms. Chemical interactions between organisms can be analyzed across all organizational levels, reaching from cell-cell interaction and intraspecific and multitrophic-level interactions to whole community interactions and environmental ecological processes. Because of their ubiquity, chemical signals that carry information (semiochemicals) can be categorized by the types of ecological interactions they mediate, such as intraspecific social communication, antagonistic interactions, and mutualism. Accordingly, this article is organized into three core areas, one formed by the chemicals mediating interactions between members of the same species (pheromones), and the others by interspecific interactions involving allomones (where the sender benefits), and synomones (where both sender and receivers benefit). A fourth group of signals, kairomones (where the receiver benefits), can comprise all other signal categories when they are perceived and utilized by a third organism that itself gains a benefit from eavesdropping on communication between others. While primary studies in chemical ecology focused on the identification of compounds mediating interactions between organisms, today’s debates are dominated by micro- and macroevolutionary aspects of chemical interactions. The very rapid growth of the chemical ecology literature over recent decades has been, in part, driven by the growing appreciation of the high economic value of understanding chemical communication, reaching from applications in pest management over the control of disease vectors in agriculture to the use of chemical signals in medicine. Moreover, the field has dramatically profited from innovations in analytical chemistry, making the separation of complex compound mixtures as well as the identification of compound structures efficient and accessible to a broader community of researchers. Recent advances in molecular ecology have aided an even more rapid mechanistic and functional analysis of semiochemicals, leading to a modern consolidation of different research fields. This collection of significant publications focuses on the functional and evolutionary analysis of chemical signals important in mediating ecological interactions. Moreover, attention has been given to publications that provide conceptual frameworks and are among the most highly cited in the respective subdisciplines. They can thus provide a good introduction for the interested reader and allow efficient forward and backward searching for more detailed information.

General Overviews

The field of chemical ecology as such is relatively young, but it has experienced a very rapid growth in the past few decades, primarily fueled by more readily available chemical analytical and molecular methods. This, on one hand, explains the limited number of concise textbooks in this field, but on the other hand, it also explains the increasing impact and explanatory power chemical ecology has in almost all fields of ecology, evolutionary biology, and biochemistry. In general, there are a number of very good summaries of the chemical ecology of particular groups of organisms, such as algae (Amsler 2008), insects (Roitberg and Isman 1992, Cardé and Miller 2004), crustaceans (Breithaupt and Thiel 2011), and vertebrates (Müller-Schwarze 2006) but a conceptional consolidation of the field of chemical ecology has rarely been undertaken. Sondheimer, et al. 1970 was one of the first comprehensive collections of studies of chemically mediated interactions by the pioneers in the field, and it was updated by another collection of studies, Eisner and Meinwald 1995. The coevolutionary aspects of chemical communication has always been a major concern of the field, and it is nicely summarized in Spencer 1988. Harborne 1993 was one of the first textbooks to reach a broader audience of students. The textbooks and collections of articles cited in this section either provide a general overview or focus on the chemical ecology of particular groups of organisms, while also allowing the extraction of the principal and generally applicable concepts.

Journals

More than 140 journals publish articles on chemical ecology, and a relatively large proportion of such articles appear in high-ranking general science journals such as Science, Nature, PNAS, PlosBio, Plos One, and Current Biology. Moreover, high-ranking ecology journals, such as Ecology Letters, Ecology, Journal of Ecology, Oecologia, Functional Ecology, and Oikos, as well as high-ranking plant science journals, such as Plant Physiology, Plant Journal, and New Phytologist, all dedicate a significant proportion of their available space to articles focusing on different aspects of chemical ecology. However, there are some journals that are specific to the field, namely the Journal of Chemical Ecology, Phytochemistry, Chemistry & Biodiversity, Molecular Ecology, and Chemoecology.

Pheromone-Mediated Interactions

With the identification of bombycol, the sexual pheromone released by the silk moth (Bombix mori) females to attract males (see Butenandt, et al. 1959), the study of the function of chemicals produced by organisms that are not used in their primary metabolism but rather in interactions with other organisms had begun. Substances that mediate interactions between members of one species were termed pheromones (Karlson and Lüscher 1959) and subcategorized by their functions in mediating specific behavioral responses (e.g., sex pheromones, alarm pheromones). Historically, the study of chemicals mediating interactions experienced its first peak with the understanding of pheromone signals mediating social interactions. Pheromones proved to be crucial in the understanding of the regulation of complex social interactions, such as those of hymenopteran ants and bees, termites, social spiders, birds, and mammals, and opened our minds to the understanding of a sensory world that proved far more important to a large number of very different groups of organisms than the relatively limited human chemosensory system made us imagine (Wyatt 2003). Chemical communication is crucial for most of the behaviors associated with cast organization, coordinated prey finding, and reproduction in most groups of insects, and is similarly crucial for the understanding of social behavior in most other animals (Wyatt 2003, Litwack 2010). Moreover, microbe social behaviors have recently been demonstrated to depend on chemical signals. A very interesting new area of chemical ecological research is the role of chemical signals mediating interactions between plants of the same or different species. Recent studies demonstrate that plants can not only respond to neighboring plant chemical signals, but that these signals may allow them to differentiate between kin and non-kin, and cause them to adjust their competition and defense phenotype accordingly.

  • Butenandt, A., R. Beckmann, D. Stamm, and E. Hecker. 1959. Über den Sexuallockstoff des Seidenspinners Bombix mori: Reindarstellung und Konstitution. Z. Naturforsch 14:283–284.

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    First original paper that reported the structural identification of a pheromone.

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  • Karlson, P., and M. Lüscher. 1959. Pheromones: A new term for a class of biologically active substances. Nature 183.4653: 55–56.

    DOI: 10.1038/183055a0Save Citation »Export Citation »E-mail Citation »

    This article provided the first thorough definition of pheromones, and so marks the beginning of the study of the chemical ecology of interactions between conspecifics.

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  • Litwack, Gerald, ed. 2010. Pheromones. Vitamins and Hormones 83. Amsterdam and Boston: Elsevier Academic.

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    Volume in a book series that includes articles from the leaders in pheromone research on all major aspects of pheromone chemical ecology, with a strong focus on insect pheromone communication.

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  • Wyatt, Tristram D. 2003. Pheromones and animal behavior: Communication by smell and taste. Cambridge, UK: Cambridge Univ. Press.

    DOI: 10.1017/CBO9780511615061Save Citation »Export Citation »E-mail Citation »

    Comprehensive textbook on pheromone ecology and biochemistry. The book summarizes knowledge on pheromone communication of many taxonomic groups, including humans, and strongly emphasizes the point that pheromone signaling can often only be understood when studied in the context of other sensory inputs.

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Invertebrate Pheromonal Communication

In insects, chemical communication is of upmost importance, and this made insects the group of organisms that allowed for the study of any type of pheromonal communication and provided the basic framework for a categorization of different types of pheromones by function (Vander Meer, et al. 1998; Cardé and Minks 1997; Sillam-Dussès 2010; Howard and Blomquist 2005). Moreover, the ongoing identification of pheromones, as well as the elucidation of their functions (Blomquist and Bagnères 2010), has allowed them to be used extensively in insect pest control, by either luring insects into traps using sex and aggregation pheromones, repelling them with alarm pheromones, or disrupting signaling by providing false pheromone sources (Mayer and McLaughlin 1990; Howse, et al. 1997; Witzgall, et al. 2010; Landolt and Phillips 1997).

  • Blomquist, Gary J., and Anne-Geneviève Bagnères, eds. 2010. Insect hydrocarbons: Biology, biochemistry, and chemical ecology. Cambridge, UK: Cambridge Univ. Press.

    DOI: 10.1017/CBO9780511711909Save Citation »Export Citation »E-mail Citation »

    The first comprehensive treatment of the study of insect cuticular hydrocarbons as chemical communication signals. The edited volume features contributions by most of the specialists in this fast-growing field, and covers almost every aspect of cuticular hydrocarbon biology, from biosynthesis to ecology.

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  • Cardé, Ring T., and Albert K. Minks, eds. 1997. Insect pheromone research: New directions. Proceedings of the First International Symposium on Insect Pheromones, held in Wageningen, The Netherlands, 6–11 March 1994. New York: Chapman & Hall.

    DOI: 10.1007/978-1-4615-6371-6Save Citation »Export Citation »E-mail Citation »

    Edited volume with contributions from most of the leading researchers on pheromone biochemistry perception and chemical ecology.

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  • Howard, Ralph W., and Gary J. Blomquist. 2005. Ecological, behavioral, and biochemical aspects of insect hydrocarbons. Annual Review of Entomology 50:371–393.

    DOI: 10.1146/annurev.ento.50.071803.130359Save Citation »Export Citation »E-mail Citation »

    Comprehensive review on arthropod chemical communication and the role of various hydrocarbons produced by arthropods in mediating interactions such as species- and gender-recognition, nestmate recognition, task-specific cues, dominance and fertility cues, chemical mimicry, and primer pheromones.

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  • Howse, Philip, Ian Stevens, and Owen Jones. 1997. Insect pheromones and their use in pest management. New York and London: Chapman & Hall.

    DOI: 10.1007/978-94-011-5344-7Save Citation »Export Citation »E-mail Citation »

    This textbook provides a broad range of examples for pheromones used for pest control. It describes all commonly used pheromone-based pest control methods, all of which are deeply based on broad knowledge of the chemical ecology of the controlled insect pests.

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  • Landolt, Peter J., and Thomas W. Phillips. 1997. Host plant influences on sex pheromone behavior of phytophagous insects. Annual Review of Entomology 42:371–391.

    DOI: 10.1146/annurev.ento.42.1.371Save Citation »Export Citation »E-mail Citation »

    Review on how sex pheromonal communication in phytophagous insects can be affected by host plant chemistry.

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  • Mayer, Marion S., and John R. McLaughlin. 1990. Handbook of insect pheromones and sex attractants. Boca Raton, FL: CRC Press.

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    Encyclopedic description of the chemical communications systems used by over 1,600 insects in sexual interactions. The volume documents in what insect species the pheromones were identified, as well as the analytical and bioassay methods used for identification and functional analysis.

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  • Sillam-Dussès, David. 2010. Trail pheromones and sex pheromones in termites. New York: Nova Science.

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    This book discusses the biology of termite pheromones and their role in mediating social interactions in termite colonies.

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  • Vander Meer, Robert K., Michael D. Breed, Karl E. Espelie, and Mark L. Winston, eds. 1998. Pheromone communication in social insects: Ants, wasps, bees, and termites. Boulder, CO: Westview.

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    Collection of articles by the leaders in the field of social insect pheromone research. Because of its thematic structure, the book can be used as both a reference and a textbook, providing detailed and basic knowledge about pheromone functions and mechanisms of action.

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  • Witzgall, Peter, Philipp Kirsch, and Alan Cork. 2010. Sex pheromones and their impact on pest management. Journal of Chemical Ecology 36.1: 80–100.

    DOI: 10.1007/s10886-009-9737-ySave Citation »Export Citation »E-mail Citation »

    Most recent and comprehensive review on the use of insect pheromone communication in agricultural pest control.

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Vertebrate Pheromone Communication

Chemical communication among members of the same species of vertebrates is increasingly well understood in respect to mechanisms of compound biosynthesis and sensorial perception (Cheal 1975, Brennan and Zufall 2006, Brennan and Kendrick 2006). However, information processing is different, potentially more complex, and less well understood than in insect study systems. For example, Wysocki and Preti 2004 reviews the literature on the difficulties in understanding human pheromonal communication. Interestingly, most insights into the function of complex vertebrate pheromone blends have been reached in the mouse (Hurst and Beynon 2004) and elephant systems (Hollister-Smith, et al. 2008; Schulte, et al. 2007). Moreover, Mason and Parker 2010 provides a valuable overview of what is known about pheromonal communication in reptiles. Most of the available literature on vertebrate pheromone communication concerns analyzing biochemistry and pheromone receptor properties, but the ecological functions of pheromonal communication (the chemical ecology) in vertebrates is certainly less well understood than in arthropods. The Chemical Signals in Vertebrates series of meeting proceedings is probably one of the best sources of information on the chemical ecology of vertebrate pheromone production (see, for example, Hurst, et al. 2006).

  • Brennan, Peter A., and Keith M. Kendrick. 2006. Mammalian social odours: attraction and individual recognition. Philosophical Transactions of the Royal Society B: Biological Sciences 361.1476: 2061–2078.

    DOI: 10.1098/rstb.2006.1931Save Citation »Export Citation »E-mail Citation »

    Review article on chemosensory properties and the chemical ecology of pheromone communication in mammals.

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  • Brennan, Peter A., and Frank Zufall. 2006. Pheromonal communication in vertebrates. Nature 444:308–315.

    DOI: 10.1038/nature05404Save Citation »Export Citation »E-mail Citation »

    Review article about transformational findings that suggest that main olfactory and vomeronasal systems have significant overlap in the chemical signals they can detect and the associated behavioral functions triggered.

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  • Cheal, Marylou. 1975. Social olfaction: A review of ontogeny of olfactory influences on vertebrate behavior. Behavioral Biology 15.1: 1–25.

    DOI: 10.1016/S0091-6773(75)92040-4Save Citation »Export Citation »E-mail Citation »

    Review article that provides a conceptional framework for the use of olfactory signals in social interactions of vertebrates, from fish to humans.

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  • Hollister-Smith, Julie A., Susan C. Alberts, and L. E. L. Rasmussen. 2008. Do male African elephants, Loxodonta africana, signal musth via urine dribbling? Animal Behaviour 76.6: 1829–1841.

    DOI: 10.1016/j.anbehav.2008.05.033Save Citation »Export Citation »E-mail Citation »

    An original study that exemplifies the ecological functional analysis of pheromone production in African elephants.

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  • Hurst, Jane L., and Robert J. Beynon. 2004. Scent wars: The chemobiology of competitive signalling in mice. Bioessays 26.12: 1288–1298.

    DOI: 10.1002/bies.20147Save Citation »Export Citation »E-mail Citation »

    Review article on the interaction between the chemical basis of mouse scents and the dynamics of their competitive scent-marking behavior. The article reviews the knowledge on the ecological function of mouse urine pheromones and contributes to the understanding of the complexity of scent signals and their information content for interacting individuals of a species.

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  • Hurst, Jane L., Robert J. Beynon, S. Craig Roberts, and Tristram D. Wyatt, eds. 2006. Chemical signals in vertebrates 11. New York: Springer Verlag.

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    Proceedings of a similarly named meeting covering research about chemical ecology, biochemistry, behavior, and neurobiology of the main olfactory and vomeronasal systems of vertebrates of a broad range of taxonomic groups. The book is part of a series of volumes that illustrates the recent history of advances in vertebrate pheromone chemical ecology.

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  • Mason, Robert T., and M. Rockwell Parker. 2010. Social behavior and pheromonal communication in reptiles. Journal of Comparative Physiology A: Neuroethology Sensory Neural and Behavioral Physiology 196.10: 729–749.

    DOI: 10.1007/s00359-010-0551-3Save Citation »Export Citation »E-mail Citation »

    Review article that focuses on pheromonal interactions in reptiles and discusses the chemical ecological aspects.

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  • Schulte, Bruce A., Elizabeth W. Freeman, Thomas E. Goodwin, Julie Hollister-Smith, and L. E. L. Rasmussen. 2007. Honest signalling through chemicals by elephants with applications for care and conservation. Applied Animal Behaviour Science 102.3: 344–363.

    DOI: 10.1016/j.applanim.2006.05.035Save Citation »Export Citation »E-mail Citation »

    Critical analysis of the sender and receiver relationship in mammal pheromonal communication, using the African elephant as an example.

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  • Wysocki, Charles J., and George Preti. 2004. Facts, fallacies, fears, and frustrations with human pheromones. Anatomical Record Part A: Discoveries in Molecular, Cellular, and Evolutionary Biology 281A.1: 1201–1211.

    DOI: 10.1002/ar.a.20125Save Citation »Export Citation »E-mail Citation »

    Review and discussion on the evidence about human pheromonal communication.

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Semiochemicals in Plant-Plant Interactions

Interactions between plants are traditionally not viewed as pheromonal interactions, because there is no nervous system processing information from conspecific neighbors. Nevertheless, new research into plant-plant interactions suggests that these types of communications are functionally equivalent to communication between animals. One early study, Baldwin and Schultz 1983, showed that plants can perceive volatile cues from neighboring plants, but this view was highly criticized at the time. Then, Karban and colleagues did a series of experiments with wild tobacco and sagebrush, demonstrating that volatile-mediated plant-plant interactions can lead to fitness consequences even between plants of different species (for review, see Karban 2008). In the early 21st century, volatile-mediated plant-plant interactions have been shown to have potentially multiple functions, the primary of which may be intra-plant signal transduction (Heil and Silva Bueno 2007, Heil and Ton 2008). Major progress has been made in understanding the mechanisms of volatile compound perception in plants (Engelberth, et al. 2004), which may also help to further reveal potential ecological functions of plant-plant signaling. For example, there is evidence that plants can differentiate between kin and non-kin, based on volatile (Karban and Shiojiri 2009) and nonvolatile (Falik, et al. 2003) chemical signals exchanged between neighbors and adjust competitive responses accordingly. Moreover, plants can perceive chemical signals from herbivore-attacked plants and ready their own defenses (Baldwin and Schultz 1983; Engelberth, et al. 2004; Karban 2008; Heil and Ton 2008).

  • Baldwin, Ian T., and Jack C. Schultz. 1983. Rapid changes in tree leaf chemistry induced by damage: Evidence for communication between plants. Science 221.4607: 277–279.

    DOI: 10.1126/science.221.4607.277Save Citation »Export Citation »E-mail Citation »

    This paper represents one of the first, and much debated, experimental tests of the hypothesis that plants can induce resistance to herbivores in response to volatile signals from neighboring plants.

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  • Engelberth, Juergen, Hans T. Alborn, Eric A. Schmelz, and James H. Tumlinson. 2004. Airborne signals prime plants against insect herbivore attack. Proceedings of the National Academy of Sciences 101.6: 1781–1785.

    DOI: 10.1073/pnas.0308037100Save Citation »Export Citation »E-mail Citation »

    First paper that demonstrates priming (the readying) of plant responses in response to volatile organic compounds emitted from herbivore-damaged neighboring plants.

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  • Falik, Omer, Perla Reides, Mordechai Gersani, and Ariel Novoplansky. 2003. Self/non-self discrimination in roots. Journal of Ecology 91.4: 525–531.

    DOI: 10.1046/j.1365-2745.2003.00795.xSave Citation »Export Citation »E-mail Citation »

    Original article that demonstrates root-signaling-mediated self/non-self recognition. Although the mechanism is not explicitly shown in the paper, the experiments suggest a potential role of chemical signals that allow differential responses to neighboring plants.

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  • Heil, Martin, and Juan Carlos Silva Bueno. 2007. Within-plant signaling by volatiles leads to induction and priming of an indirect plant defense in nature. Proceedings of the National Academy of Sciences 104.13: 5467–5472.

    DOI: 10.1073/pnas.0610266104Save Citation »Export Citation »E-mail Citation »

    Original paper that demonstrates within-plant signal transduction of information about the damage status by means of herbivore-induced volatile organic compound emissions with effects on whole plant resistance.

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  • Heil, Martin, and Jurriaan Ton. 2008. Long-distance signalling in plant defence. Trends in Plant Science 13.6: 264–272.

    DOI: 10.1016/j.tplants.2008.03.005Save Citation »Export Citation »E-mail Citation »

    Review article that focuses on the crucial role of plant volatile organic compounds in transporting information from damaged to undamaged tissues within and between plants. The paper reviews evidence for plant-plant communication induced by both herbivores and pathogens, and discusses the functional significance of volatile-mediated plant-plant interactions.

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  • Karban, Richard. 2008. Plant behaviour and communication. Ecology Letters 11.7: 727–739.

    DOI: 10.1111/j.1461-0248.2008.01183.xSave Citation »Export Citation »E-mail Citation »

    Review on plant communication phenomena and their ecological consequences, including anticipation of future environmental conditions, priming of responses, memory and trans-generational induced resistance.

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  • Karban, Richard, and K. Shiojiri. 2009. Self-recognition affects plant communication and defense. Ecology Letters 12.6: 502–506.

    DOI: 10.1111/j.1461-0248.2009.01313.xSave Citation »Export Citation »E-mail Citation »

    Original study that demonstrates the herbivore-induced volatile organic compounds can function as kin-recognition cues and result in differential induced responses in kin and non-kin neighbors.

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Allomonal Communication

Another major theme of chemical ecology is the chemistry of defenses. Many organisms produce compounds that are not needed for their primary physiological functions or for interactions with members of their own species, but are merely produced to defend against attackers. Well known are extremely potent toxins such as those of reptiles (Weldon, et al. 2008, cited under Vertebrate Chemical Defenses) and amphibians (Daly 1995, also under Vertebrate Chemical Defenses), unpleasant repellent compounds of many mammals (Koh, et al. 2009 and Burger 2005, both cited under Vertebrate Chemical Defenses) such as skunks (Wood, et al. 2002, under Vertebrate Chemical Defenses) and opossums, but also the vast diversity of toxic compounds produced by plants and insects (Berenbaum 1995). The latter two (plants and insects) became major model systems in chemical ecology because they include the largest part of biodiversity on the planet, and because of their easier approachability as study systems. Moreover, their species diversity is matched by the diversity of interactions they are engaged in with each other. There is a plethora of herbivorous insect species that attack and consume plants, and an equally large number of insect species that have complex mutualistic relationships with plants. Thus, the study of the chemical ecology of plant-insect interactions resulted in the formulation of major theory and, as a consequence, the fastest growth in publication output in the field in recent years. Berenbaum 1995 illustrates how all major and fundamental principles in chemical ecology can be addressed by studying plant-insect interactions.

Invertebrate Chemical Defenses

Defensive compounds secreted by arthropods are as diverse as this most species-rich group of higher organisms (Blum 1981, Aldrich 1988, Eisner and Meinwald 1966). In fact, it has been hypothesized that the arthropod “phyletic dominance” could have originated in large part because of the evolution of chemical defenses (Meinwald and Eisner 1995). Indeed, arthropod defensive chemicals are, in general, multifunctional and toxic, repellent, insecticidal, antimicrobial, and surfactant, with a large array of potential target organisms (Dettner 1987; Eisner, et al. 2005). Some groups of arthropods are characterized by their diverse defensive chemistries, such as beetles (Dettner 1987) and Heteroptera (Aldrich 1988). Moreover, the compounds can be biosynthesized by the arthropods themselves (Eisner and Meinwald 1966), or they can be sequestered from food resources (Duffey 1980, Blum 1981). While many compounds have been identified, their defensive function (e.g., the increased fitness of individuals that produce certain compounds, as compared to individuals not able to produce those compounds)—and thus their chemical ecology—is often difficult to study, which limits the number of studies that have investigated arthropod chemical defenses in a population and community ecology context (Pasteels, et al. 1983). Nevertheless, most of the reviews in this collection provide frameworks of hypotheses that identify research trajectories beyond the identification of compounds. In addition to insects, a strong focus of chemical ecological research was put on marine invertebrates beginning in the early 1990s. Lindquist and Hay 1996 provides a valuable review on the defense chemistry of marine invertebrate larvae.

  • Aldrich, Jeffrey R. 1988. Chemical ecology of the Heteroptera. Annual Review of Entomology 33:211–238.

    DOI: 10.1146/annurev.en.33.010188.001235Save Citation »Export Citation »E-mail Citation »

    Comprehensive treatment of the chemical ecology of a taxonomic group (Heteroptera) that is rich in defensive compound production. The paper provides a nice example of the diversity of compounds produced by a group of organisms and their multiple ecological functions.

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  • Blum, Murray S. 1981. Chemical defenses of arthropods. New York: Academic Press.

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    Monograph that represents the first comprehensive treatment of arthropod chemical ecology covering all allomone compound classes known to be biosyntheiszed or sequestered (from food) by arthropods for their own defense. Detailed descriptions of chemical defenses are provided for centipedes, scorpions, spiders, millipedes, and insects.

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  • Dettner, Konrad. 1987. Chemosystematics and evolution of beetle chemical defenses. Annual Review of Entomology 32:17–48.

    DOI: 10.1146/annurev.en.32.010187.000313Save Citation »Export Citation »E-mail Citation »

    This review uses a phylogenetic approach to categorize beetle defense chemistry. It provides a valuable example of a conceptual framework for studying arthropod defense chemical ecology.

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  • Duffey, Sean S. 1980. Sequestration of plant natural products by insects. Annual Review of Entomology 25:447.

    DOI: 10.1146/annurev.en.25.010180.002311Save Citation »Export Citation »E-mail Citation »

    This review provides a comprehensive overview of insect sequestration of plant chemicals in general, and takes a special focus on the defensive function of the sequestered compounds. It also summarizes the factors that contribute to sequestration.

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  • Eisner, Thomas, Maria Eisner, and Melody V. S. Siegler. 2005. Secret weapons: Defenses of insects, spiders, scorpions and other many-legged creatures. Cambridge, MA: Belknap Press of Harvard Univ. Press.

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    Excellent book that describes selected examples of the chemical ecology of arthropod defenses, including the chemistry, physiology, behavioral biology, ecology, and evolutionary biology of defense compound production.

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  • Eisner, Thomas, and Jerrold Meinwald. 1966. Defensive secretions of arthropods. Science 153.3742: 1341–1350.

    DOI: 10.1126/science.153.3742.1341Save Citation »Export Citation »E-mail Citation »

    This review is a classic in arthropod defense chemical ecology. It provides an overview of the compound classes (known at the time) that are used by arthropods for defense. Moreover, the work provides a conceptual framework for the study of arthropod allomones.

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  • Lindquist, Neils, and Mark E. Hay. 1996. Palatability and chemical defense of marine invertebrate larvae. Ecological Monographs 66.4: 431–450.

    DOI: 10.2307/2963489Save Citation »Export Citation »E-mail Citation »

    Comprehensive review on defenses strategies of marine invertebrate larvae. Although this work is not focused on defensive chemistry, it positions the chemical ecology of the discussed organisms in an integrative framework of hypotheses on when and how organisms cope with predators under certain environmental conditions.

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  • Meinwald, Jerrold, and Thomas Eisner. 1995. The chemistry of phyletic dominance. Proceedings of the National Academy of Sciences 92:14–18.

    DOI: 10.1073/pnas.92.1.14Save Citation »Export Citation »E-mail Citation »

    Review and concept article that states the hypothesis that the evolution of chemical defenses was a prerequisite for arthropods to become the phyletically most dominating group of higher organisms on earth.

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  • Pasteels, J. M., J.-C. Grégoire, and M. Rowell-Rahier. 1983. The chemical ecology of defense in arthropods. Annual Review of Entomology 28:263–289.

    DOI: 10.1146/annurev.en.28.010183.001403Save Citation »Export Citation »E-mail Citation »

    Key review that discusses insect defense compound production from a macro- and microevolutionary perspective. Moreover, it reviews knowledge on target specificity of allomones, the biological significance of compound mixtures, and intraspecific variation in defense compound production.

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Vertebrate Chemical Defenses

Vertebrate defensive chemistry is not as well studied as that of insects and plants, but it provides some striking examples of complex and efficient defense strategies. Prominent examples come from amphibians that use very toxic skin secretions to defend themselves against attackers (Daly 1995, Hanifin 2010). Similarly, reptiles—in particular snakes—are well-known for their potent venoms. Less is known, however, about reptile integument secretions. Weldon, et al. 2008 provides a review on the growing literature on this subject. In addition to reptiles and amphibians, mammal chemical defenses have been relatively well studied (Burger 2005). Well-known examples in this area, such as skunk defensive sprays (Wood, et al. 2002) and the recently identified complex platypus venoms (Koh, et al. 2009), are notable and provide nice case studies for a mechanistic and functional analysis of defensive compound production. More recently, and in general in chemical ecology, the study of the structure-function relationships of compounds within compound classes common to multiple taxa has become an important research direction. Wong, et al. 2007 provides a review of such a structure-function analysis of a compound class—defensins—that is widely distributed among vertebrate taxa.

  • Burger, B. V. 2005. Mammalian semiochemicals. In Chemistry of Pheromones and Other Semiochemicals II. Edited by S. Schulz, 231–278. Berlin: Springer-Verlag Berlin.

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    Review article on the chemical characterization and functional analysis of urine, anal gland secretions, and exocrine gland secretions of mammals. The review includes hormonal functions but also discussion of potential defensive functions of mammal semiochemicals.

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  • Daly, John W. 1995. The chemistry of poisons in amphibian skin. Proceedings of the National Academy of Sciences 92:9–13.

    DOI: 10.1073/pnas.92.1.9Save Citation »Export Citation »E-mail Citation »

    Review on amphibian skin chemistry, including a categorization of compounds into those biosynthesized by the amphibians and those that are sequestered from food resources. The relative role of sequestration and chemical alteration by microorganisms of compounds is discussed and provides an excellent framework of hypotheses for the study of the chemical ecology of amphibian defense chemistry.

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  • Hanifin, Charles T. 2010. The chemical and evolutionary ecology of tetrodotoxin (TTX) toxicity in terrestrial vertebrates. Marine Drugs 8.3: 577–593.

    DOI: 10.3390/md8030577Save Citation »Export Citation »E-mail Citation »

    The review describes the chemicals and evolutionary ecology of one of the most well-known animal toxins in the world, tetrodotoxin, in amphibians rather than in marine organisms, where the toxin is widely distributed among a broad range of taxa.

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  • Koh, Jennifer M. S., Paramjit S. Bansal, Allan M. Torres, and Philip. W. Kuchel. 2009. Platypus venom: source of novel compounds. Australian Journal of Zoology 57.4: 203–210.

    DOI: 10.1071/ZO09040Save Citation »Export Citation »E-mail Citation »

    Review on the compound-rich venomous secretions of the platypus, one of two mammal species producing venom. The paper characterizes the chemistry and discusses the chemical ecology and the potential application of the compounds in medicine.

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  • Weldon, Paul J., Birte Flachsbarth, and Stefan Schulz. 2008. Natural products from the integument of nonavian reptiles. Natural Product Reports 25:738–756.

    DOI: 10.1039/b509854hSave Citation »Export Citation »E-mail Citation »

    This paper reviews the epidermal and glandular chemistry of nonavian reptiles and discusses their functions. The authors organized the paper taxonomically and collected data from more than 170 references.

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  • Wong, Jack H., Lixin Xia, and T. B. Ng. 2007. A review of defensins of diverse origins. Current Protein & Peptide Science 8.5: 446–459.

    DOI: 10.2174/138920307782411446Save Citation »Export Citation »E-mail Citation »

    Review of defensin proteins and their antimicrobial functions in a broad range of taxonomic groups.

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  • Wood, William F., Brian G. Sollers, Gwen A. Dragoo, and Jerry W. Dragoo. 2002. Volatile components in defensive spray of the hooded skunk, Mephitis macroura. Journal of Chemical Ecology 28.9: 1865–1870.

    DOI: 10.1023/A:1020573404341Save Citation »Export Citation »E-mail Citation »

    This research paper provides a nice example of the analysis of mammal allelochemicals, in this case the volatile components of the defensive spray of the hooded skunk. The paper reviews all the literature on skunk defensive sprays.

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Plant Chemical Defenses

Plants are probably the best-studied group of organisms in chemical ecology. Much of the theory about the mechanisms and functions of defense trait production, particularly chemicals, has been derived from plant systems, mostly in their interactions with arthropods. In many cases the biosynthesis and ecological function of plant secondary metabolite production is well understood, and modern genetic and analytical tools help to further the study of micro- and macroevolutionary aspects of plant defenses. A coevolutionary arms race of chemical defense and counter-defenses was first proposed for plant-insect interactions in the 1960s (see Ehrlich and Raven 1964), and this generated a rapid growth in publication output in the field of chemical ecology. Thus, the publications featured here can only hint at the extraordinarily rich body of knowledge existing on the ecology and evolution of plant defenses. The chemical ecology of plant insect interactions as a field has been most influenced by questions about coevolutionary interactions between plants and their attackers (Rosenthal and Berenbaum 1992), the functionality of diversity and redundancy in defense compound production (Rosenthal and Berenbaum 1991, Rosenthal and Berenbaum 1992), the phenotypic plasticity (induced production) of defense chemistry production and its ecological consequences (Karban and Baldwin 1997), and the understanding of genetic and transcriptional regulation of plant secondary metabolite production (Kessler and Baldwin 2002). In addition, the question of the physiological and ecological costs of plant defenses was central to the development of an evolutionary hypothetical framework for the understanding of plant-herbivore coevolution. In the debate about this question, and in further developing plant defense theory, some publications stand out. One is Herms and Mattson 1992, which outlines the concept of plant-defense production being traded off with the investment of resources into other life functions. Another one, Coley and Barone 1996, provides a comprehensive and seminal review on different aspects of plant defense theory, using tropical plant-herbivore systems as examples. Hay and Fenical 1988 combines the well-studied community ecology aspects of marine plant–herbivore interactions with chemical ecology. Walters 2010 is one of the first comprehensive textbooks on plant defenses against herbivores and pathogens.

  • Coley, P. D., and J. A. Barone. 1996. Herbivory and plant defenses in tropical forests. Annual Review of Ecology and Systematics 27.1: 305–335.

    DOI: 10.1146/annurev.ecolsys.27.1.305Save Citation »Export Citation »E-mail Citation »

    Seminal review on the ecological and evolutionary consequence of plant-insect interactions in tropical systems. It provides a general overview over multiple aspects of plant defense theory and a road map for future research in chemical ecology.

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  • Ehrlich, Paul R., and Peter H. Raven. 1964. Butterflies and plants: A study in coevolution. Evolution 18.4: 586–608.

    DOI: 10.2307/2406212Save Citation »Export Citation »E-mail Citation »

    This is a seminal paper that like no other influenced thinking and research in the study of the chemical ecology of plant-herbivore interactions. It explains and collects supportive data on the hypothesis that plants and insect herbivores are engaged in a coevolutionary arms race of evolution of toxic defenses in the plant and counter-defenses in the herbivore. The paper has significantly influenced theory in chemical ecology in general

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  • Hay, Mark E., and William Fenical. 1988. Marine plant-herbivore interactions: The ecology of chemical defense. Annual Review of Ecology and Systematics 19.1: 111–145.

    DOI: 10.1146/annurev.es.19.110188.000551Save Citation »Export Citation »E-mail Citation »

    This review article applies plant defense theory to marine plant–herbivore interaction systems and discusses the chemical ecology of marine plants.

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  • Herms, Daniel A., and William J. Mattson. 1992. The dilemma of plants: To grow or defend. Quarterly Review of Biology 67.3: 283–335.

    DOI: 10.1086/417659Save Citation »Export Citation »E-mail Citation »

    This is one of the most cited papers in the field discussing the growth-defense hypothesis. This paper and the described hypothesis have sparked research in a fundamental proportion of the field and continue to be influential in the study of the chemical ecology of plant defenses and in the study of plant-herbivore interactions in general.

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  • Karban, Richard, and Ian T. Baldwin. 1997. Induced responses to herbivory. Chicago: Univ. of Chicago Press.

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    Textbook that summarizes theory and data on plant-induced responses to herbivory. The book provides a solid base from which to explore the field of plant phenotypic plasticity in resistance trait expression. It also summarizes all major theories surrounding plant chemical defenses in general.

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  • Kessler, André, and Ian T. Baldwin. 2002. Plant responses to insect herbivory: The emerging molecular analysis. Annual Review of Plant Biology 53:299–328.

    DOI: 10.1146/annurev.arplant.53.100301.135207Save Citation »Export Citation »E-mail Citation »

    Review of the literature on mechanisms and ecological consequences of induced plant responses to herbivory. The paper provides a comprehensive overview over different induction mechanisms of plant responses and discusses ecological and evolutionary aspects of induced direct and indirect resistance in the plant defense theory framework.

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  • Rosenthal, G., and M. Berenbaum, eds. 1991. Herbivores: Their interactions with secondary plant metabolites. Vol. 1, The chemical participants. 2d ed. San Diego, CA: Academic Press.

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    This first of two volumes (see Rosenthal and Berenbaum 1992) that provide a comprehensive review of chemical compound classes that are involved in plant defenses and discusses the mechanisms of their production and function. First edition, edited by Gerald A. Rosenthal and Daniel H. Janzen, published in 1979.

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  • Rosenthal, G., and M. Berenbaum, eds. 1992. Herbivores: Their interactions with secondary plant metabolites. Vol. 2, Ecology and evolutionary processes. San Diego, CA: Academic Press.

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    This second of two volumes (see Rosenthal and Berenbaum 1991) complements the first one and reviews the functional and evolutionary aspects of the chemical ecology of plant defenses against herbivores. It includes all the most cited theories surrounding plant chemical defenses and efficiently reviews the relevant literature. Both books provide a valuable source of information and innovation for the experienced and inexperienced reader in the field.

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  • Walters, Dale. 2010. Plant defense: Warding off attack by pathogens, herbivores and parasitic plants. Chichester, UK: Wiley-Blackwell.

    DOI: 10.1002/9781444328547Save Citation »Export Citation »E-mail Citation »

    Textbook that gives a broad and valuable overview over plant defenses mechanisms and the ecological consequences of plant defenses against herbivores and pathogens. The book is thought to provide an entry for the newcomer to the field and a reference for the experienced reader.

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Synomonal Communication

Symbiosis, the living together of organisms of different species, is highly regulated and, in most cases, relies on a complex web of chemical signals. It does not matter if the interactions are antagonistic, requiring chemical signals that keep the antagonist away or allow the attacker to overcome its host’s defenses, or if the interactions are mutualistic, where the chemical signals allow coordinated metabolic or behavioral adjustments of the interacting organisms (Schaefer and Ruxton 2011). Antagonistic and mutualistic interactions can thus be viewed as two opposite extremes of symbiotic interactions. Mutualistic interactions are also often mediated by chemical signals (synomones), allowing a coordinated optimization of interspecific interactions, and ultimately resulting in the reciprocal benefit of the interacting species. Again, most of the examples for chemically mediated mutualistic interactions come from plant-insect systems. A well-known interaction between plants and insects is that of plants with their pollinators, the chemistry of which is only beginning to be understood (Raguso 2008). Moreover, many plant species respond to herbivore damage with an increased production of volatile organic compounds, which can function as cues for host/prey to search out natural enemies of the herbivores, such as predators and parasitoids (Vet and Dicke 1992). In these cases the plant-derived chemical signals facilitate host/prey search behavior of the natural enemies, to the benefit of both the natural enemy and the plant (Dicke and van Loon 2000). The discovery of plant volatile organic compound–mediated indirect resistance in agricultural (Turlings, et al. 1990) and natural systems (Kessler and Baldwin 2001) had a strong influence in the field and resulted in a new integrated view on plant secondary metabolite production in the plant defenses context. Many, if not most, plant secondary metabolites can thus be expected to have multiple functions, mediating both antagonistic and mutualistic interactions with other organisms (Gershenzon and Dudareva 2007). A nice example for multifunctionality is root sesquiterpenes that mediate the mutualistic interaction between plants and arbuscular mycorrhizal fungi (Akiyama, et al. 2005), but that are also utilized by antagonists. Chemical signaling is likely a property common to most mutualistic interactions and is fundamental to mutualisms that involve insects. One of the most striking examples of such chemistry-mediated mutualisms is that between Lycanidae butterflies and their tending ants. Axén, et al. 1996 provides a case study on the chemical ecology of this system.

  • Akiyama, Kohki, Ken-ichi Matsuzaki, and Hideo Hayashi. 2005. Plant sesquiterpenes induce hyphal branching in arbuscular mycorrhizal fungi. Nature 435:824–827.

    DOI: 10.1038/nature03608Save Citation »Export Citation »E-mail Citation »

    Original study that demonstrates a crucial role of plant-derived and root-exuded sesquiterpenes (strigolactones) in mediating the mutualistic interaction between plant roots and mycorrhizal fungi.

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  •  Axén, Annkristen H., Olof Leimar, and Veronika Hoffman. 1996. Signalling in a mutualistic interaction. Animal Behaviour 52.2:321–333.

    DOI: 10.1006/anbe.1996.0178Save Citation »Export Citation »E-mail Citation »

    Case study of the mutualistic interaction between Lycanidae butterfly larvae and their mutualistic ants, which is mediated by chemical signals.

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  • Dicke, Marcel, and Joop J. A. van Loon. 2000. Multitrophic effects of herbivore-induced plant volatiles in an evolutionary context. Entomologia Experimentalis Et Applicata 97:237–249.

    DOI: 10.1046/j.1570-7458.2000.00736.xSave Citation »Export Citation »E-mail Citation »

    Review of the literature on herbivore-induced volatile organic compound emissions and their role in attracting natural enemies (predators and parasitoids) of herbivores. The potential fitness effects of attracting natural enemies of herbivores for the plant are critically discussed. The authors conclude that the attraction of natural enemies is a chemically mediated mutualistic interaction between plants and insects.

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  • Gershenzon, Jonathan, and Natalia Dudareva. 2007. The function of terpene natural products in the natural world. Nature Chemical Biology 3:408–414.

    DOI: 10.1038/nchembio.2007.5Save Citation »Export Citation »E-mail Citation »

    This work comprehensively reviews the ecological functions of plant terpenoid production, which mediates myriad interactions of plants with other organisms, a big proportion of which are mutualistic.

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  • Kessler, André, and Ian T. Baldwin. 2001. Defensive function of herbivore-induced plant volatile emissions in nature. Science 291:2141–2144.

    DOI: 10.1126/science.291.5511.2141Save Citation »Export Citation »E-mail Citation »

    First evidence that indirect plant defenses by means of herbivore-induced volatile organic compound emission is mediating mutualistic interactions between plants and predators in a wild study system.

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  • Raguso, Robert A. 2008. Wake up and smell the roses: The ecology and evolution of floral scent. Annual Review of Ecology, Evolution, and Systematics 39:549–569.

    DOI: 10.1146/annurev.ecolsys.38.091206.095601Save Citation »Export Citation »E-mail Citation »

    Critical review on the role of floral scents in mediating plant-pollinator interactions. The paper discusses the ecology and evolutionary biology of private channels of unusual compounds, unique ratios of more widespread compounds, or multicomponent floral filters as ways that scent can promote specialization in plant-pollinator relationships.

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  • Schaefer, H. Martin, and Graeme T. Ruxton. 2011. Plant-animal communication. Oxford and New York: Oxford Univ. Press.

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    This textbook provides a theoretical framework for the evolutionary ecology of plant-animal communication, with a strong emphasis on mutualistic interactions. It includes plant interactions with herbivores, pollinators, seed dispersers and predators of plant herbivores. While all types of communication are discussed, there is a strong emphasis on chemical communication.

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  • Turlings, T. C. J., J. H. Tumlinson, and W. J. Lewis. 1990. Exploitation of herbivore-induced plant odors by host-seeking parasitic wasps. Science 250.4985: 1251–1253.

    DOI: 10.1126/science.250.4985.1251Save Citation »Export Citation »E-mail Citation »

    One of the first and most influential studies demonstrating that herbivore-induced volatile organic compounds can attract natural enemies of herbivores, such as hymenopteran parasitoids, to the plant, and so function as indirect defense.

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  • Vet, Louise E. M., and Marcel Dicke. 1992. Ecology of infochemical use by natural enemies in a tritrophic context. Annual Review of Entomology 37:141–172.

    DOI: 10.1146/annurev.en.37.010192.001041Save Citation »Export Citation »E-mail Citation »

    One of the first comprehensive reviews on plants’ ability to attract natural enemies of their herbivores by means of herbivory-induced volatile organic compound emission. The work discusses ecological and evolutionary aspects of the herbivore-induced emission of volatiles.

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The Chemistry of Cheating and Lying

All chemical signals produced by an organism can potentially be utilized by another organism. This includes the attraction of antagonists by pheromones or synomones that have evolved to mediate positive interactions. But it also includes the production of compounds by one species that mimic signals of another species, allowing deception about the presence of mating partners, breeding sites, or food, or allowing a species to be chemically camouflaged (Dettner and Liepert 1994). The study of chemical cheating and lying is very new and promises to be one of the most productive research fields in chemical ecology (Vereecken and McNeil 2010). The literature on the subject is currently dominated by food- and sex-deceptive plant-pollinator systems, and by examples of chemical mimicry/camouflage of insects that are food parasites in nests of social insects. Ayasse, et al. 2003 and Brodman, et al. 2009 provide excellent case studies for sex-deceptive and bee alarm pheromone-mediated attraction, respectively, of pollinators to orchid flowers. Deceptive pollination in orchids provides some of the most striking examples of chemical cheating and lying and is increasingly being studied on an evolutionary level. Jersáková, et al. 2006 critically reviews the evolution of deceptive pollination and includes a large number of additional examples for the phenomenon. In comparison to sex deception, which is largely restricted to orchids, food and brood-site deception seems to be more widely distributed in the plant kingdom. Urru, et al. 2011 reviews the chemical ecology of brood-site deception, with a discussion of evolutionary and ecological aspects. There are a number of interesting studies on insects hijacking the communicational systems of social and nonsocial Hymenoptera. For example, Hafernik and Saul-Gershenz 2000 describes the chemical ecology of blister beetles mimicking female bee pheromones to get into their nests, where they are food parasites. Similarly, Nash, et al. 2008 shows that larvae blue butterflies mimic the cuticular hydrocarbons of their ant hosts.

  • Ayasse, Manfred, Florian P. Schiestl, Hannes F. Paulus, Fernando Ibarra, and Wittko Francke. 2003. Pollinator attraction in a sexually deceptive orchid by means of unconventional chemicals. Proceedings of the Royal Society B: Biological Sciences 270.1514: 517–522.

    DOI: 10.1098/rspb.2002.2271Save Citation »Export Citation »E-mail Citation »

    This study represents the chemical ecology of one of the prime examples of sexually deceptive orchids. Orchid flowers mimic sex pheromones of the pollinating bees, and thus attract pollinators without presenting an award.

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  • Brodmann, Jennifer, Robert Twele, Wittko Francke, Luo Yi-bo, Song Xi-qiang, and Manfred Ayasse. 2009. Orchid mimics honey bee alarm pheromone in order to attract hornets for pollination. Current Biology 19.16: 1368–1372.

    DOI: 10.1016/j.cub.2009.06.067Save Citation »Export Citation »E-mail Citation »

    Original study on an orchid that mimics bee alarm pheromone to attract a bee predator for pollination.

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  • Dettner, Konrad, and Caroline Liepert. 1994. Chemical mimicry and camouflage. Annual Review of Entomology 39:129–154.

    DOI: 10.1146/annurev.en.39.010194.001021Save Citation »Export Citation »E-mail Citation »

    This review summarizes examples of two very important ways of coping with antagonists, chemical mimicry and camouflage. The paper covers themes like the integration into colonies of social insects, the exploitation of mutualistic interactions, exploitation of brood care of solitary insects, luring of prey, reproduction, chemical interrelationships between plants and insects, and Müllerian mimicry of warning odors, all mediated by secondary metabolites produced by arthropods.

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  • Hafernik, John, and Leslie Saul-Gershenz. 2000. Beetle larvae cooperate to mimic bees. Nature 405:35–36.

    DOI: 10.1038/35011129Save Citation »Export Citation »E-mail Citation »

    Study that reports the collaboration between nonsocial blister beetles to chemically mimic the presence of a bee female for male bee attraction. Male bees collect the blister beetle larvae and transfer them to female bees, which carry them into the nest.

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  • Jersáková, Jana, Steven D. Johnson, and Pavel Kindlmann. 2006. Mechanisms and evolution of deceptive pollination in orchids. Biological Reviews 81.2: 219–235.

    DOI: 10.1017/S1464793105006986Save Citation »Export Citation »E-mail Citation »

    Critical review on the ecology and evolutionary biology of deceptive orchids, including a discussion of the chemical ecology of the mimicry.

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  • Nash, David R., Thomas D. Als, Roland Maile, Graeme R. Jones, and Jacobus J. Boomsma. 2008. A mosaic of chemical coevolution in a large blue butterfly. Science 319.5859: 88–90.

    DOI: 10.1126/science.1149180Save Citation »Export Citation »E-mail Citation »

    Case study on chemical mimicry using larvae of blue butterflies mimicking the cuticular hydrocarbons of their ant hosts as an example.

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  • Urru, Isabella, Marcus C. Stensmyr, and Bill S. Hansson. 2011. Pollination by brood-site deception. Phytochemistry 72.3: 1655–1666.

    DOI: 10.1016/j.phytochem.2011.02.014Save Citation »Export Citation »E-mail Citation »

    Review on the chemical ecology of brood-site deception, with a discussion of the ecology and evolutionary biology of the underlying interactions.

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  • Vereecken, N. J., and J. N. McNeil. 2010. Cheaters and liars: Chemical mimicry at its finest. Canadian Journal of Zoology 88.7: 725–752.

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    Critical review of chemical mimicry, with a particular emphasis on cases relating to pollination and obtaining food resources. The paper provides a conceptional framework for the study of chemical mimicry.

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Future Perspectives and Applications

Recent advances in chemical ecology have been strongly influenced by developments in analytical chemistry and the merge with molecular biology techniques (Degenhardt, et al. 2003, Seybold 2004). These advances also promise an increased application of chemical-ecological knowledge (Schoonhoven, et al. 2005). So far most applications concern the control of insect pests in agriculture and the control of insect vectors of diseases. Chemical-ecological principles have found their place in integrated pest management (see Koul and Cuperus 2007), and have themselves been used as alternative strategies for pest control. For example, insect pheromones are now widely used to monitor and control insect pests in agricultural fields (Reddy and Guerrero 2004). A review on the management of the codling moth in Witzgall, et al. 2008 nicely illustrated how important the knowledge of the chemical ecology of the system is in order to successfully control pests. In addition to controlling insect pests, weeds can be efficiently controlled with allelopathic compounds from other plants (Anaya 1999). Probably the most successful and most influential application of chemical ecology in pest control is the push-pull strategy that was developed to control insect pests and weeds in East African corn fields (Cook, et al. 2007). It is not only agricultural pests that can be controlled by means of chemical ecology, however. Logan and Birkett 2007 provides a comprehensive review on the control of blood-sucking insects using chemical-ecological knowledge of their systems.

  • Anaya, Ana Luisa. 1999. Allelopathy as a tool in the management of biotic resources in agroecosystems. Critical Reviews in Plant Sciences 18.6: 697–739.

    DOI: 10.1016/S0735-2689(99)00397-4Save Citation »Export Citation »E-mail Citation »

    A transformational review that proposes the use of allelopathic compound production in plants for weed control in agriculture. The literature on allelopathy as well as on traditional practices is reviewed.

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  • Cook, Samantha M., Zayaur R. Khan, and John A. Pickett. 2007. The use of push-pull strategies in integrated pest management. Annual Review of Entomology 52:375–400.

    DOI: 10.1146/annurev.ento.52.110405.091407Save Citation »Export Citation »E-mail Citation »

    Review on one of the most successful practical applications of chemical ecological knowledge in agricultural pest control, with a very important outlook on the potential of methods, such as the push-pull agricultural practice, for the development of a more sustainable agriculture.

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  • Degenhardt, Joerg, Jonathan Gershenzon, Ian T. Baldwin, and André Kessler. 2003. Attracting friends to feast on foes: Engineering terpene emission to make crop plants more attractive to herbivore enemies. Current Opinion in Biotechnology 14.2: 169–176.

    DOI: 10.1016/S0958-1669(03)00025-9Save Citation »Export Citation »E-mail Citation »

    This review article describes the potential of combining modern chemical and molecular ecology knowledge with biotechnology tools to find new ways of pest control in agriculture.

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  • Koul, Opender, and Gerrit W. Cuperus, eds. 2007. Ecologically based integrated pest management. Wallingford, UK: CABI.

    DOI: 10.1079/9781845930646.0000Save Citation »Export Citation »E-mail Citation »

    This is a collection of review chapters on integrated pest management. Although not focused on the application of chemical ecology, the book illustrates the broad application of chemical ecological knowledge when utilizing natural defenses on crop plants and the attraction of predators, parasitoids, and entomopathogens in agricultural pest control.

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  • Logan, James G., and Michael A. Birkett. 2007. Semiochemicals for biting fly control: Their identification and exploitation. Pest Management Science 63.7: 647–657.

    DOI: 10.1002/ps.1408Save Citation »Export Citation »E-mail Citation »

    Review article on the control of blood-sucking insects with methods derived from chemical ecological research.

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  • Reddy, Gadi V. P., and Angel Guerrero. 2004. Interactions of insect pheromones and plant semiochemicals. Trends in Plant Science 9.5: 253–261.

    DOI: 10.1016/j.tplants.2004.03.009Save Citation »Export Citation »E-mail Citation »

    Review article on the chemical ecology of insect pheromones and plant volatiles as the two most important volatile semiochemicals in insect behavior, and on their applications in pest control.

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  • Schoonhoven, Louis M., Joop J. A. van Loon, and Marcel Dicke. 2005. Insect-plant biology. 2d ed. Oxford: Oxford Univ. Press.

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    This book provides an up-to-date synthesis of the rapidly expanding field of chemical ecology of plant-insect interactions. It provides a review and broad outlook on the potential applications of chemical ecological knowledge on plant-insect interactions in agricultural pest control.

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  • Seybold, Steven J. 2004. Preface: The eighth day of discovery: Molecular biology comes to chemical ecology. Journal of Chemical Ecology 30.12: 2327–2333.

    DOI: 10.1007/s10886-004-7939-xSave Citation »Export Citation »E-mail Citation »

    Preface to a special issue in the Journal of Chemical Ecology about the transforming merge between chemical ecology and molecular biology. The papers in the issue discuss future perspectives for the field and applications of chemical ecological knowledge.

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  • Witzgall, Peter, Lukasz Stelinski, Larry Gut, and Don Thomson. 2008. Codling moth management and chemical ecology. Annual Review of Entomology 53:503–522.

    DOI: 10.1146/annurev.ento.53.103106.093323Save Citation »Export Citation »E-mail Citation »

    Review article on one of the most successful examples of using insect pheromones as a mating disruption agent in the biological control of a major apple pest.

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LAST MODIFIED: 05/23/2012

DOI: 10.1093/OBO/9780199830060-0023

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